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An Investigation of the Floral Morphogenesis of Bonatea Speciosa (Orchidaceae)

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An Investigation of the Floral Morphogenesis of Bonatea Speciosa (Orchidaceae) S. Afr.1. Bot., 1989, 55(4): 433--437 433 An investigation of the floral morphogenesis of Bonatea speciosa (Orchidaceae) H. Kurzweil Bolus Herbarium, University of Cape Town, Rondebosch, 7700 Republic of South Africa The flower morphology and ontogeny of Bonatea speciosa are described, and special attention is paid to the early development of the gynostemium. The lateral appendages of the latter ('auricles') are shown to be compound structures, consisting of a filament appendage and a small bulge likely to be a staminode. The median carpel apex is an enormous hood-like structure and lacks a receptive portion, the receptive stigma being entirely confined to the apices of the stigma stalks which correspond to the lateral carpel apices. All important stages are documented by SEM micrographs. Die blommorfologie en ontogenie van Bonatea speciosa word beskryf en spesiale aandag word aan die vroee ontwikkeling van die ginostegium gegee. Die laterale aanhangsels van laasgenoemde ('aurikels') is saamge­ stelde strukture, bestaande uit 'n filament en 'n klein geswolle dee I wat waarskynlik 'n staminodium is. Die mediaan-vrugblaarpunt is 'n groot kappievormige struktuur waarvan die ontvanklike dee I ontbreek, die ontvanklike stempel is geheel en al beperk tot die punte van die stempelsteeltjies wat met die laterale vrug­ blaarpunte ooreenstem. Aile belangrike stadiums word deur aftaselektronmikrograwe gestaaf. Keywords: Bonatea speciosa, floral morphogenesis, Orchidaceae, SEM investigation Introduction Materials and Methods Bonatea is a comparatively small genus of 20 species Flowers and floral buds were collected in the Cape which is distributed from Arabia through East Africa to Peninsula near Cape Town (L1andudno, Kurzwei! HK Cape Town in South Africa with the majority of species 859: voucher in BOL), and immediately fixed and found in South Africa's summer-rainfall area (Stewart preserved in 'Kew Cocktail' (H2 0 :alcohol:formalde­ et. ai. 1982). The plants grow terrestrially in grassland , hyde:glycerine = 15.3:16:1:1). The material was dehy­ forest and coastal shrub. The genus is generally placed in drated and critical-point dried after the technique of the tribe Orchideae of the subfamily Orchidoideae on Gerstenberger & Leins (1978) . Mounted samples were account of having a gynostemium with an erect basi to­ sputter-coated with Au/Pd and observed and photo­ nous anther (Dressler 1981). graphed with a Cambridge Stereoscan S-200 scanning The floral structures, in particular the curious long electron microscope at 5 and 10 k V. basal processes of the thecae and the stalked stigmas, are very similar to those in the related genus Habenaria , in Observations which genus Bonatea was also included in the past. It differs by the basal fusion of the lip , the anterior petal The most striking features of the flower are the white, lobes and the lateral sepals to the stigma stalks and by sharply projecting, stalked stigmas; the lateral rostellum the occurrence of a tooth-like lip appendage in front of lobes attached to the long, narrow anther canals; and the the spur entrance. However, morphological comparisons white, twisted anterior petal lobes (Figure 1). The lateral with Habenaria were not made in this study, as no sepals, the anterior petal lobes and the lip are basally detailed information on Habenaria is available. A few fused to the stigma stalks (which can be recognized by data on the flower morphology and ontogeny of Haben­ the broken tissue on the lower surface of the latter; aria species were presented by Vermeulen (1959) and arrowhead in Figure 2). Kurzweil (1987). The green posterior lobes of the petals form , together The structure of the adult flower of B. speciosa (L.f.) with the green, deeply galeate median sepal , a hood Willd. was studied by Trimen (1864) , Vermeulen (1959) enclosing the gynostemium. The green and slightly and Weale (1867). However, no ontogenetic information oblique lateral sepals are unlobed in all ontogenetic is available in order to properly analyze and interpret the stages. structure of flowers in this genus. In the open flower, the two petal lobes are very similar The main aim of the paper is to examine if, and in how in size and general shape (Figure 12d). However, the far, the development of the curious odd-shaped gynoste­ posterior lobe represents the original petal, the anterior mium architecture of Bonatea is basically similar to the lobe developing from a small basal process on the ontogeny described in some species of Orchideae - already existing posterior petal lobe (Figure 12d- f). Orchidinae with their comparatively simple gynoste­ Interestingly, the petal also has a small posterior process mium structure (Kurzweil 1987) and, in particular, if the in early stages of development (arrowhead in Figure ontogeny of the auricles of Bonatea conforms to what 12f) . was described in the above-mentioned paper. B. speciosa The green lip is deeply three-lobed and is variously was chosen for an investigation of the floral development twisted and contorted. The side lobes develop much as material of this species is easily available in Cape later than the midlobe (Figure 12a-c), the initial lip Town. being completely unlobed. A slender spur is inserted on 434 S.-Afr.Tydskr. Plantk., 1989, 55(4) the lip base and is initiated in middle ontogenetic stages. elongate and angled upwards in this species, and are The small lip appendage in front of the spur entrance is attached to the lateral rostellum arms (Figure 10 & 11). developed in late stages. The anther is thus typically 'basitonous' (i .e . the viscidia The anther of the gynostemium is erect from the early are attached to the bases of the thecae). As in related developmental stages onward. Initially it is ovate and genera, the division of the anther into thecae develops slightly exceeds the perianth lobes in length (Figure 4) , early (Figure 4). The short, rounded apical connective but develops into an elongate structure distinctly shorter process develops from the apex of the young anther, than the tepals (Figures 5-11). In the early ontogenetic which was not divided by the gap that separates the stages the thecae are slightly narrower in their basal thecae (Figures 5, 6 & 10) . parts. These develop gradually into the slender anther The gynostemium has lateral appendages which are canals found in the mature flower, which are strongly commonly termed 'auricles' (au + b in Figures 2 & 3) . Figures 1-3 Flower and gynostemium in the mature stage. 1. Flower in front view. 2. Gynostemium in side view. 3. Gynostemium in front view. A I = anther, au = auricle, b = basal bulge, I = lip , PI = petal, ro = rostellum , Sl = median sepal, S3 = lateral sepal, stg = stigma, vi = viscidium (the abbreviations for the anther and the perianth lobes with their subscript numbers are those commonly used in orchid literature). Figure 1: bar = 10 mm; Figures 2- 3: bar = 1 mm. S.Afr.J. Bot., 1989, 55(4) 435 Figures 4-11 Various stages of the gynostemium development. 4-6. Gynostemium in an early stage in front view. 7-9. Early development of auricles and basal bulges. 10. Gynostemium in front view. 11. Base of gynostemium in a late stage. AI = anther, au = auricle, b = basal bulge, Ie = lateral carpel apices, mc = median carpel apex, P2 = petal. SEM micrographs. Figures 4-9: bars = 0.1 mm; Figures 10-11 : bars = 1 mm. 436 S.-Afr.Tydskr. Plantk. , 1989,55(4) Discussion sp The ontogeny of the perianth lobes is almost identical to that of other Orchidoideae, which was described in an earlier paper (Kurzweil 1987). However, the occurrence of a minute posterior petal lobe in early stages is noteworthy, and possibly indicates a tendency towards three-lobing of the inner tepals - a sequence which is clearly expressed in the inner median tepal (= 'lip'). Whereas three-lobing of the lip is very common in Orchi­ deae, three-lobing of petals is apparently rather rare but occurs also in some Habenaria species (Kurzweil, in b prep.). a The sequence of organ initiation and their initial shape in the early development of the gynostemium also proved to be very similar to that of related species of the tribe Orchideae (Kurzweil 1987). The anther of Bonatea speciosa is erect as in all species al of the tribe Orchideae, distinguishing them from the Diseae (Disinae, Coryciinae, Satyriinae) with mostly reflexed anthers. Long, narrow anther canals are rather common in the subtribe Habenariinae in which Bonatea is generally placed, but are rare in the Orchidinae. The auricles on the anther base of this species are compound structures and originate from both dorsolat­ eral anther processes and small bulges initiated on the receptacle at the base of the petals (here termed 'basal bulges'). The dorsolateral anther processes are not homologous to staminal structures as they are initiated d e f on the sides of the already existing anther. The bulges on Figure 12 Development of lip (a-c) and petal (d-f). al the receptacle are homologous with the 'basal bulges' anterior petal lobe, sp = spur. Bars = 1 mm. described in Kurzweil (1987) , which, due to their early initiation and their place of origin are probably lateral staminodes of the inner staminal whorl. This mode of They are made up of two structures of different origin: development of the auricles (double origin) corresponds whereas the posterior and larger portion develops from a well with the auricle development observed in other dorsolateral anther appendage (au in Figures 7-9), the Orchideae (Kurzweil 1987) . However, the auricles as a anterior portion originates from the receptacle between whole were in the past referred to as staminodes by most the petal base and the gynostemium (here termed 'basal authors, as their dual origin has not been noted before. bulge' ; b in Figures 7-9).
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